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. Author manuscript; available in PMC: 2022 May 26.
Published in final edited form as: Annu Rev Plant Biol. 2022 Feb 15;73:149–172. doi: 10.1146/annurev-arplant-070221-050044

Table 1.

Histone variant genes, proteins, and functions in Arabidopsis thaliana

Histone variants Arabidopsis Genes ArabidopsisLoci Protein Chaperones/remodelers Function Knockout/knockdown Arabidopsis phenotype Modification
General development Stress responses
H3.3 HTR4
HTR5
HTR8
At4g40030
At4g40040
At5g10980
H3.3
H3.3
H3.3
HIRA (95)
ATRX(33)
Transcriptional activation (123, 127, 144, 156) Flowering time(156);
Male Gametogenesis (54)); Post-germination development (5); Cell proliferation and organogenesis (101)
Regulate the expression of responsive and hypervariable genes(145) htr5/htr8, are phenotypically normal ; Triple mutants cause lethality; htr4/htr4;htr8/htr8 plants that carried either amiR-HTR5-I or amiR-HTR5-II (h3.3kd) exhibit leaf serration, incomplete sterility, and growth defects (145) H3.3K36me3 (89)
H2A.Z HTA8
HTA9
HTA11
At2g38810
At1g52740
At3g54560
H2A.Z.8
H2A.Z.9
H2A.Z.11
SWR1 complex (42, 137) transcriptional activation and repression Flowering time (16, 29, 86, 98, 124); vegetative to reproductive phase transition (42, 148); Inflorescence architecture (11); Germline development (157); Circadian clock (133) Temperature response (24, 66, 132);
Phosphate deficiency (40, 126, 151) Drought stress (11, 128);
Immunity (6);
Salt stress (93)
Double/ triple mutants show developmental aberrations (83, 94); Triple mutants are viable but show reduced fertility (20);
Knockdown of all three H2AZ genes cause early flowering (16
H2A.Z acetylation (25); H2A.Z monoubiquitination (H2A.Zub) (44); H2A.Z SUMOylation and methylation?
H2A.X HTA3
HTA5
At1g54690
At1g08880
H2A.X.3
H2A.X.5
FACT? (45) DNA damage response (13, 67, 112); transcription activation (146) H2A.X is unessential for Arabidopsis development (50) genotoxic stress(141) The single/ double mutants are viable, fertile and indistinguishable from wild type (50) phosphorylated H2A.X (67, 112)
H2A.W HTA6
HTA7
HTA12
At5g59870
At5g27670
At5g02560
H2A.W.6
H2A.W.7
H2A.W.12
DDM1 (100) Chromatin condensation (150) H2A.W is unessential for Arabidopsis development
(10)
Not determined H2A.W triple mutants are indistinguishable from the wild type (10 phosphorylated H2A.W (75, 112)
H1 HON1
HON2
HON3
At1g06760
At2g30620
At2g18050
H1.1
H1.2
H1.3
NRP1 and NRP2?(91, 102) Chromatin condensation (14, 114) Flowering time,
Seed dormancy,
Lateral root, Stomata and callus development (114)
male and female gametogenesis (120)
Drought stress (2)
Combined light and water deficiency (116)
Triple mutants are viable but show extended dormancy, early flowering, increased root density and lateral root numbers, and altered stomata pattern (114) phosphorylation, acetylation, mono- and dimethylation, formylation, ,crotonylation and propionylation (64)